Female Preference Intensities on Different Call Characteristics and Symmetry of Preference Above and Below the Mean in the Iberian Midwife Toad Alytes cisternasii

The intensity of female preference for call characteristics is measured in two‐speaker playback tests with female Iberian midwife toads ‘Alytes cisternasii’ using an experimental design that allows quantitative comparisons across characters. Female preferences do not appear to be symmetrical above and below the population mean, female preference being more efficient at discriminating against low quality competing males that at selecting optimum males among good ones. Female preference is significantly more effective selecting males calling at higher call rates (a highly variable call characteristic probably related to the physical state of the male) rather than selecting calls of low dominant frequency (a characteristic of low intra‐individual variation, related to male size and age).     

Acoustic Competition in Male Midwife Toads Alytes obstetricans and Alytes cisternasii: Response to Neighbor Size and Calling Rate. Implications for Female Choice

Effects of neighbor on male calling behavior was studied through playback experiments of synthetic calls to males of two species of midwife toads. The responses of resident males were scored considering two temporal parameters (call duration and calling rate) and one spectral parameter (dominant frequency). The sounds used for the playback tests included two levels of fundamental frequency (correlated with male size) and two levels of call repetition rate. In both species, resident males only changed their calling rate in the presence of an intruder, and the response was different for synthetic calls with two levels of dominant frequencies and with two calling rates. Resident size was not significantly correlated with the magnitude of the change in the calling rate. On the other hand, resident calling rate was significantly and positively correlated with the magnitude of the increase in calling rate of the stimulus. The maximum relative increase in calling rate was observed in A. cisternasii. In phonotaxis tests, females are preferentially attracted to calls emitted at a higher rate confirming the importance of changes in calling rate for female attraction.     

Microsatellite markers for the Mallorcan midwife toad Alytes muletensis

The highly endangered midwife toad Alytes muletensis occurs only on the Mediterranean island of Mallorca. Little is known about genetic variability and dispersal among the 19 breeding sites (six of which stem from reintroductions) of this amphibian. The value of such information should be high for successful management of this species. We report here the characterization of eight polymorphic microsatellite loci in A. muletensis using individuals from a single population.     

Reproductive stage and history affect the phonotactic preferences of female midwife toads, Alytes muletensis

We investigated whether the reproductive stage or reproductive history of females affects their preferences for male calls that differ in frequency. We repeatedly tested gravid female midwife toads prior to and immediately after mating, and during ovulation. When females were ovulating they were much more consistent in their choice of call than when they were gravid but not ovulating. When females had recently mated they still showed positive phonotaxis, but did not reliably discriminate between alternative calls. This study is the first to test gravid female anurans repeatedly at different stages in their reproductive cycle (ovulating or not; pre- and postmating). It highlights the need to treat with caution the results from population-based preference studies in which individuals are tested only once, and in which variation in reproductive state is not taken into account. Copyright 2000 The Association for the Study of Animal Behaviour.     

Low gene flow but high genetic diversity in the threatened Mallorcan midwife toad Alytes muletensis

We investigated fine-scale genetic structuring in the rare and vulnerable Mallorcan midwife toad Alytes muletensis using eight polymorphic microsatellite markers. The current range of this amphibian is restricted to some 19 sites of which six are derived from reintroductions, all located in the mountain ranges of Mallorca. We sampled tadpoles from 14 pools covering 10 natural sites and two reintroduction sites for microsatellite DNA analyses. Relatively high levels of genetic variation were found in most pools (H(E) = 0.38-0.71, allelic richness = 2.6-6.2). Only at one pool has the population recently gone through a bottleneck. Dispersal between pools in different torrents does not occur whereas downstream dispersal between pools within the same torrent does happen at low frequencies. This occasional exchange of individuals does not lead to neighbouring pools in the same torrent being panmictic. This can be concluded because all F(ST) values (0.12-0.53) differ significantly from zero and STRUCTURE analyses identified neighbouring pools as separate populations. Furthermore, assignment and migration tests showed little exchange between neighbouring pools. If upstream locations or complete torrents go extinct, they are unlikely to be recolonized naturally. For conservation purposes, reintroductions of tadpoles to sites where local extinctions have occurred may therefore be advisable.     

Male-male Competition and Female Mate Choice through Courtship Display in the Territorial Damselfish Stegastes nigricans

Both sexes of the herbivorous damselfish Stegastes nigricans maintain individual feeding territories. These territories are distributed contiguously, forming distinct colonies. Females visit male territories to spawn, and eggs are guarded by males until hatching. Male-male competition and female mate choice were studied in two colonies of different size compositions. Only larger individuals bred in both colonies. Some males in the large colony, that were larger than the breeding males in the small colony, did not succeed in reproducing probably because of severe attacks by the larger males while courting. However, females did not choose large size among breeding males. The most important male characteristic in female choice was the frequency of courtship displays in both colonies. Females in the large colony chose males mainly on the basis of the frequency of displays conducted in the females' territories, whereas females in the small colony chose males on the basis of the frequency of displays conducted in the males' territories. This difference may be a result of the difference in colony size. The distances between females' and males' territories were much greater in the large colony, and, because females cannot see courtship displays conducted in distant male territories, males in the large colony may have had to visit female territories frequently in order to conduct courtship near the females.     

Within- and Among-Population Variation in Chytridiomycosis-Induced Mortality in the Toad Alytes obstetricans

Background

Chytridiomycosis is a fungal disease linked to local and global extinctions of amphibians. Susceptibility to chytridiomycosis varies greatly between amphibian species, but little is known about between- and within-population variability. However, this kind of variability is the basis for the evolution of tolerance and resistance evolution to disease.

Methodology/Principal Findings

In a common garden experiment, we measured mortality after metamorphosis of Alytes obstetricans naturally infected with Batrachochytrium dendrobatidis. Mortality rates differed significantly among populations and ranged from 27 to 90%. Within populations, mortality strongly depended on mass at and time through metamorphosis.

Conclusions/Significance

Although we cannot rule out that the differences observed resulted from differences in skin microbiota, different pathogen strains or environmental effects experienced by the host or the pathogen prior to the start of the experiment, we argue that genetic differences between populations are a likely source of at least part of this variation. To our knowledge, this is the first study showing differences in survival between and within populations under constant laboratory conditions. Assuming that some of this intraspecific variation has a genetic basis, this may suggest that there is the potential for the evolution of resistance or tolerance, which might allow population persistence.     

Female preference in complex acoustical environments in the midwife toads Alytes obstetricans and Alytes cisternasii

Gravid females of Alytes obstricans and Alytes cisternasii weretested with synthetic calls in seven-speaker playback tests.A first, "mean-centered" test presented calls with frequenciesrepresenting an array of different calls spanning over the rangeof the population (±2.25 SD). In this test, females ofboth species approached a synthetic call that was lower thanthe average call frequency of the male population although thedifference was significant only for A. obstetricans. The regressionbetween female weight and size and preferred frequency was notsignificant in either species. These results confirm the reportedtrends of females preferring lower frequency calls (correspondingto larger males) based on two-speaker playback tests for A.obstetricans. For A. cisternasii, the lack of significance ofthe seven-speaker test suggests that the preference trend previouslyfound in two-speaker tests may be obscured in more complex acousticalenvironments. A second "supernormal stimulus" test presentedfemales with calls ranging from the lowest frequency valuesof the male population (–2.25 SD) and lower, up to –6.75SD beyond the range. In both species females preferentiallyapproached calls higher than the mean frequency of the stimuluspresented. This result suggests that in both cases selectionfor low frequencies is not open ended, and that the preferredfrequency is within the range of the male population.     

Evidence for the Introduction of Lethal Chytridiomycosis Affecting Wild Betic Midwife Toads (Alytes dickhilleni)

Batrachochytrium dendrobatidis is an unpredictable pathogen for European amphibian species, and existing field surveillance studies likely underestimate the scope of its distribution and effects. Mass mortality episodes recorded in Europe indicate that investigations of unstudied species should focus on members of the frog family Alytidae. Here, we report the combined results of a field survey and laboratory observations of field collected Alytes dickhilleni. Our data support the hypothesis that B. dendrobatidis has recently emerged in at least two disjunct locations in the species range and populations across much of the species range lack evidence of infection pathogen. Tadpoles taken into the laboratory from sites with infection experienced 70% mortality, unlike those taken into the laboratory from uninfected sites, and both infection and strength of infection was associated with mortality in animals collected from infected locations. Several conservation interventions are underway in response to our study, including the establishment of a captive assurance colony, a public awareness campaign, and experimental tests of disease mitigation schemes.     

Call Intensity and Female Preferences in the European Green Toad

In the present paper we study the pattern of variation in call intensity in a natural population of the European green toad (Bufo viridis), and we analyse females preferences for this property by means of playback experiments. Although call sound pressure level (SPL) shows little within‐bout variation, we found significant positive correlation between call SPL and fundamental frequency: on average, an increase of 6 dB SPL produces a 100‐Hz increase in fundamental frequency. When females were given a choice between two calls differing by 10 and 6 dB SPL, they significantly preferred the loudest call, whereas they did not discriminate between calls differing by 3 dB. To test whether a 3‐dB difference reflects a sensory or behavioural limit, we carried out three experiments where females were given a choice between two calls differing in both duration (4 s and 6 s, respectively) and SPL (3 dB). In all three experiments, the longer call attracted a larger number of females, but the 3‐dB difference did not show any significant effect. Finally, we investigated the relationships between call intensity and frequency on female preferences. Previous experiments showed size‐dependent preferences for calls with lower‐than‐average frequencies (1.3 kHz) over calls with higher‐than‐average frequencies (1.6 kHz). Here, we show that this weak preference is abruptly reversed when the highest‐pitched call is broadcast at an intensity 6 dB higher than the alternative.     

Risky Courtship: Background Contrast,Ornamentation, and Display Behavior of Wolf Spiders Affect Visual Detection by Toad Predators

Males that search widely for females and perform conspicuous courtship displays run a high risk of being detected by their predators. Therefore, gains in reproductive success might be offset by increased mortality due to predation. Male brush‐legged wolf spiders (Schizocosa ocreata) with larger decorative traits (foreleg tufts) are preferred by females as mates, but are more readily detected by predators. However, predation risk may also be influenced by the interaction between components of signals and the environment in which signaling occurs. Courting male spiders were readily accepted as prey by a sympatric predator, the American toad (Anaxyrus americanus). We used video playback to tease apart the interactive effect between visual signals and the signaling environment on the ability of toads to detect courting spiders as a function of distance, background contrast, the presence or absence of male foreleg tufts, and behavioral activity. The response of toads to video sequences of male spiders was similar to their response to live male spiders. Toad response varied over distance toward spiders displayed against high contrast (sunny) vs. low contrast (shaded) backgrounds. Beyond 30 cm, more toads detected courting male spiders against light, ‘sunny’ backgrounds and detected them faster when compared to the same spider stimulus against darker, ‘shady’ backgrounds. In choice tests, toads oriented more often toward courting males with leg tufts than those without. Toad responses also varied with male spider behavior in that only videos of moving males were attacked. Latency to orient and detection by toads was significantly greater for walking males than courting males, and this effect was most evident at distances between 30 cm and 50 cm. Results supported that courting wolf spiders are at significant risk of predation by visually acute predators. Distance, background contrast, and the presence of foreleg decorations influence detection probability. Thus, the same complex visual signals that make males conspicuous and are preferred by females can make males more vulnerable as prey to toads.     

Courtship Behavior and Detection of Female Receptivity in the Parasitoid Wasp Urolepis rufipes

Once a Urolepis rufipes male mounted, the female beat her antennae against his mouth and clypeus. Immediately after he swept his antennae rapidly downward and extruded his mouthparts, her abdomen rose as she opened her genital orifice. Almost simultaneously he backed up for copulation and she folded her antennae against her head. Neither her abdomen rising nor her antennal folding were essential to his backing up as determined from their timing and from experiments in which her abdomen was sealed or her antennae were removed. Females did not open their genital orifice if with a sealed-mouth male; and antennae-removed females did not open even in the few cases where untreated males extruded their mouthparts. Unlike a closely related species, females mounted by sealed-mouth males did not open in response to air from containers of mating pairs.     

Female Genitalia Concealment Promotes Intimate Male Courtship in a Water Strider

Violent coercive mating initiation is typical for animals with sexual conflict over mating. In these species, the coevolutionary arms-race between female defenses against coercive mating and male counter-adaptations for increased mating success leads to coevolutionary chases of male and female traits that influence the mating. It has been controversial whether one of the sexes can evolve traits that allow them to “win” this arms race. Here, we use morphological analysis (traditional and scanning electron micrographs), laboratory experiments and comparative methods to show how females of a species characterized by typical coercive mating initiation appear to “win” a particular stage of the sexual conflict by evolving morphology to hide their genitalia from direct, forceful access by males. In an apparent response to the female morphological adaptation, males of this species added to their typically violent coercive mounting of the female new post-mounting, pre-copulatory courtship signals produced by tapping the water''s surface with the mid-legs. These courtship signals are intimate in the sense that they are aimed at the female, on whom the male is already mounted. Females respond to the signals by exposing their hidden genitalia for copulatory intromission. Our results indicate that the apparent victory of coevolutionary arms race by one sex in terms of morphology may trigger evolution of a behavioral phenotype in the opposite sex.     

The Effect of Male Competition on the Courtship Song of Drosophila melanogaster

The courtship song of Drosophila has been extensively used as a model system for studies of sexual selection and species recognition. Traditionally, the courtship song has been recorded from males placed individually with a female. However, under natural conditions females are exposed to multiple courting males, and the effect of their joint signal on mate recognition by the female is not yet understood. Here, we recorded the courtship behavior of D. melanogaster males singing either individually to a female lpar;1:1) or in the presence of an additional male lpar;2:1). We compared the structure of the male song in the two experimental designs. Our results show that when two males courted a female their songs could overlap each other. Males produced a significantly different signal in the presence of competition; the duration of each song component was significantly shorter and the rate of singing was markedly lower. The present study demonstrates that male competition can dramatically alter the acoustic signal detected by the female.     

Male Courtship Pheromones Affect Female Behaviour in the Swordtail Characin (Corynopoma riisei)

Pheromones constitute an important cue used by both males and females during courtship. Here, we investigate the effect of male pheromones on female behaviour in the swordtail characin (Corynopoma riisei), a species of fish where males have a caudal pheromone gland which has been suggested to affect female behaviour during courtship. We subjected female C. riisei to male courtship pheromones and investigated the effect on both female behaviour and brain serotonergic activity levels compared to a control group. While no difference in serotonergic activity was found, the pheromone‐treated females showed lower stress levels compared to the control group. Furthermore, pheromone‐treated females increased locomotor activity over time, while a decrease in locomotor activity was observed in the control group. These results suggest that the male courtship pheromones may serve to reduce female stress and increase female activity, possibly to aid males in gaining access to females and facilitating sperm transfer.     

视觉能触发沟渠豹蛛雄蛛对雌蛛的求偶行为

以沟渠豹蛛Pardosa laura成熟雄蛛的求偶延迟时间和触肢伸展次数作为参数,观察研究了雄蛛通过视觉与处女雌蛛拖丝单独和共同诱导情况下的求偶反应,发现视觉是触发雄蛛求偶行为的主要信号,而雌蛛的拖丝则不能单独激发雄蛛的求偶行为,表明雄蛛能够通过视觉接受雌蛛的信号。在此基础上,又测定了雄蛛通过视觉接受亚成体雌蛛、处女雌蛛和交配未产卵雌蛛信号时的求偶反应,发现雄蛛对处女雌蛛和交配未产卵雌蛛的求偶动作次数显著多于亚成体雌蛛,表明雄蛛能够通过视觉区分雌蛛是否成熟,但是仅依靠视觉无法判断雌蛛是否已经交配。本实验表明视觉在沟渠豹蛛雄蛛对雌蛛识别过程中起着非常重要的作用,再次证实狼蛛的视觉比较发达。     

Male Courtship Pheromones Induce Cloacal Gaping in Female Newts (Salamandridae)

Pheromones are an important component of sexual communication in courting salamanders, but the number of species in which their use has been demonstrated with behavioral evidence remains limited. Here we developed a behavioral assay for demonstrating courtship pheromone use in the aquatically courting Iberian ribbed newt Pleurodeles waltl. By performing an in-depth study of the courtship behavior, we show that females invariably open their cloaca (cloacal gaping) before engaging in pinwheel behavior, the circling movement that is the prelude to spermatophore uptake. In contrast, cloacal gaping was not observed in failed courtships, where females escaped or displayed thanatosis. Since gaping mainly occurred during male amplexus and cloacal imposition, which is the obvious period of pheromone transfer, we next investigated whether male courtship water (i.e., water holding courtship pheromones) alone was able to induce this reaction in females. These tests showed that courtship water induced cloacal gaping significantly more than water, even in the absence of a male. Cloacal gaping thus provides a simple and robust test for demonstrating courtship pheromone use in the Iberian ribbed newt. Since opening the cloaca is an essential prerequisite for spermatophore pick-up in all internally fertilizing salamanders, we hypothesize that variations on this assay will also be useful in several other species.